Supplementation with sterols improves food quality of a ciliate for Daphnia magna

Experimental results provide evidence that trophic interactions between ciliates and Daphnia are constrained by the comparatively low food quality of ciliates. The dietary sterol content is a crucial factor in determining food quality for Daphnia. Ciliates, however, presumably do not synthesize sterols de novo. We hypothesized that ciliates are nutritionally inadequate because of their lack of sterols and tested this hypothesis in growth experiments with Daphnia magna and the ciliate Colpidium campylum. The lipid content of the ciliate was altered by allowing them to feed on fluorescently labeled albumin beads supplemented with different sterols. Ciliates that preyed upon a sterol-free diet (bacteria) did not contain any sterols, and growth of D. magna on these ciliates was poor. Supplementation of the ciliates' food source with different sterols led to the incorporation of the supplemented sterols into the ciliates' cells and to enhanced somatic growth of D. magna. Sterol limitation was thereby identified as the major constraint of ciliate food quality for Daphnia. Furthermore, by supplementation of sterols unsuitable for supporting Daphnia growth, we provide evidence that ciliates as intermediary grazers biochemically upgrade unsuitable dietary sterols to sterols appropriate to meet the physiological demands of Daphnia.     

The influence of bacteria-dominated diets on Daphnia magna somatic growth, reproduction, and lipid composition

We explored how dietary bacteria affect the life history traits and biochemical composition of Daphnia magna, using three bacteria taxa with very different lipid composition. Our objectives were to (1) examine whether and how bacteria-dominated diets affect Daphnia survival, growth, and fecundity, (2) see whether bacteria-specific fatty acid (FA) biomarkers accrued in Daphnia lipids, and (3) explore the quantitative relationship between bacteria availability in Daphnia diets and the amounts of bacterial FA in their lipids. Daphnia were fed monospecific and mixed diets of heterotrophic (Micrococcus luteus) or methanotrophic bacteria (Methylomonas methanica and Methylosinus trichosporium) and two phytoplankton species (Cryptomonas ozolinii and Scenedesmus obliquus). Daphnia neonates fed pure bacteria diets died after 6-12?days and produced no viable offspring, whereas those fed pure phytoplankton diets had high survival, growth, and reproduction success. Daphnia fed a mixed diet with 80% M.?luteus and 20% of either phytoplankton had high somatic growth, but low reproduction. Conversely, Daphnia fed mixed diets including 80% of either methane-oxidizing bacteria and 20% Cryptomonas had high reproduction rates, but low somatic growth. All Daphnia fed mixed bacteria and phytoplankton diets had strong evidence of both bacteria- and phytoplankton-specific FA biomarkers in their lipids. FA mixing model calculations indicated that Daphnia that received 80% of their carbon from bacteria assimilated 46?±?25% of their FA from this source. A bacteria-phytoplankton gradient experiment showed a strong positive correlation between the proportions of the bacterial FA in the Daphnia and their diet, indicating that bacterial utilization can be traced in this keystone consumer using FA biomarkers.     

Food quality of mixed bacteria–algae diets for Daphnia magna

Bacteria can comprise a large fraction of seston in aquatic ecosystems and can therefore significantly contribute to diets of filter-feeding zooplankton. To assess the effect of three heterotrophic bacteria (Flavobacterium sp., Pseudomonas sp. and Escherichia coli) on survival, growth and egg production of juvenile Daphnia magna during six-day growth experiments, five ratios of bacteria–Scenedesmus obliquus mixtures were fed. Potential growth-limiting effects mediated by essential biochemicals were assessed upon supplementation of pure bacterial diets with a sterol (cholesterol) or a polyunsaturated fatty acid (EPA). Pure bacterial diets always had detrimental effects on Daphnia. However, cholesterol supplementation of Flavobacterium sp. enhanced growth rates of Daphnia. Diets containing Pseudomonas impaired Daphnia growth even at low dietary proportions (20%), indicating their toxicity. In contrast, Daphnia grew at relative high dietary proportions of Flavobacterium sp. and E. coli (80–50%). In fact, diets containing small proportions of these heterotrophic bacteria (Flavobacterium ≤50%, E. coli 20%) even significantly increased Daphnia growth rates compared to pure algal diets, indicating a nutritional upgrading by these bacteria. Our results suggest that the relative contribution of bacteria and phytoplankton to total dietary carbon as well as their phylogenetic composition strongly influence Daphnia fitness and potentially other filter-feeding zooplankton under field conditions.     

Essential fatty acid content and the phosphorus to carbon ratio in cultured algae as indicators of food quality for Daphnia

1. A series of experiments examined growth of Daphnia magna on three algal diets ( Rhodomonas minuta , Scenedesmus acutus and Synechococcus sp.) at varying physiological states [nitrogen and phosphorus (P) limitations] to test whether variation in algal fatty acid and/or elemental composition can predict Daphnia growth.
2. These algae differed widely in their essential fatty acid (EFA) composition while phosphorus (P) or nitrogen limitation had only a small influence on their ω3-polyunsaturated fatty acid (PUFA) content.
3. Individually, algal ω3-PUFA content explained 69% of the variation in the normalised growth of D. magna, while algal phosphorus content explained 11% of the variation. Quantitative models for D. magna growth used both algal ω3-PUFA content and algal C : P ratio as food quality indices. Together, algal ω3-PUFA content and C : P ratio explained 70% of the variation in the normalised growth rate of D. magna .
4. Our results indicate that EFA influenced algal food quality much more strongly than P. The EFA and mineral P impacts appear to be independent.     

Variability in protist grazing and growth on different marine Synechococcus isolates

Grazing mortality of the marine phytoplankton Synechococcus is dominated by planktonic protists, yet rates of consumption and factors regulating grazer-Synechococcus interactions are poorly understood. One aspect of predator-prey interactions for which little is known are the mechanisms by which Synechococcus avoids or resists predation and, in turn, how this relates to the ability of Synechococcus to support growth of protist grazer populations. Grazing experiments conducted with the raptorial dinoflagellate Oxyrrhis marina and phylogenetically diverse Synechococcus isolates (strains WH8102, CC9605, CC9311, and CC9902) revealed marked differences in grazing rates-specifically that WH8102 was grazed at significantly lower rates than all other isolates. Additional experiments using the heterotrophic nanoflagellate Goniomonas pacifica and the filter-feeding tintinnid ciliate Eutintinnis sp. revealed that this pattern in grazing susceptibility among the isolates transcended feeding guilds and grazer taxon. Synechococcus cell size, elemental ratios, and motility were not able to explain differences in grazing rates, indicating that other features play a primary role in grazing resistance. Growth of heterotrophic protists was poorly coupled to prey ingestion and was influenced by the strain of Synechococcus being consumed. Although Synechococcus was generally a poor-quality food source, it tended to support higher growth and survival of G. pacifica and O. marina relative to Eutintinnis sp., indicating that suitability of Synechococcus varies among grazer taxa and may be a more suitable food source for the smaller protist grazers. This work has developed tractable model systems for further studies of grazer-Synechococcus interactions in marine microbial food webs.     

The relative importance of algae and bacteria as food for Daphnia longispina (Cladocera) in a polyhumic lake

SUMMARY. 1. Grazing on algae and bacteria by the planktonic cladoceran, Daphnia longispina , was studied in a small polyhumic lake with low phytoplankton primary production in southern Finland.
2. D. longispina filtered algae at average rates of 0.09-0.82 ml ind⁻¹ h⁻¹. The filtering rates on bacteria were 26-92% of those on algae in parallel experiments.
3. From June to August algae, including mixotrophic and heterotrophic forms, comprised 56-93% of the food ingested by D. longispina. In mid-September and early October, when the Daphnia population was declining and the algal biomass was low, bacteria comprised 73% and 55% respectively of the food of Daphnia.
4. For D. longispina , an energy pathway via bacteria and bactivorous flagellates is probably a more important link to allochthonous organic matter than direct utilization of epilimnetic bacteria.     

Food quantity and quality regulation of trophic transfer between primary producers and a keystone grazer (Daphnia) in pelagic freshwater food webs

The transfer of energy and nutrients from plants to animals is a key process in all ecosystems. In lakes, inefficient transfer of primary producer derived energy can result in low animal growth rates, accumulation of nuisance phytoplankton blooms and dissipation of energy from the ecosystem. Most research on carbon transfer efficiency in pelagic food webs has focused on either food quantity or food quality, with the latter considered separately as either elemental stoichiometry or biochemical composition. The natural occurrence and magnitude of these types of growth limitations and their combined effects on Daphnia , a keystone grazer in pelagic freshwater ecosystems, are largely unknown. Our empirical models predict that the strength and nature of food quantity and quality limitation varies greatly with lake trophic state (total phosphorus, TP) and that Daphnia growth rates and thus energy and nutrient transfer efficiency are highest in lakes with intermediate trophic status (TP 10–25 μg l⁻¹). We predict that food availability place the greatest constraint on Daphnia growth in nutrient poor lakes (TP≤4 μg l⁻¹). Phosphorus limitation of Daphnia growth increased with decreasing TP, but the overall effect was never predicted to be the dominant constraining factor. Eicosapentaenoic acid (EPA, 20:5ω3) limitation was predicted to occur in both nutrient poor and nutrient rich lakes and placed the primary constraint on food quality in the most productive lakes. Two contrasting EPA-models gave different results on the magnitude of EPA-limitation, implying that additional food quality factors decrease Daphnia growth at high TP. In conclusion, the model predicts that Daphnia growth should peak in mesotrophic lakes, food quantity will place the greatest constraint on growth in oligotrophic lakes and EPA will primarily limit growth in eutrophic lakes.     

Seasonal dynamics of crustacean zooplankton, heterotrophic nanoflagellates and bacteria in a shallow, eutrophic lake

1. The seasonal development of crustacean zooplankton, heterotrophic nanoflagellates (HNF) and bacteria was examined in Grosser Binnensee, a shallow, eutrophic lake in northern Germany. The grazing impact of Daphnia on bacteria and nanoflagellates was estimated from field data on population abundances and from clearance rates obtained in laboratory experiments. 2. The seasonal succession of zooplankton showed distinct peaks of Daphnia magna, cyclopopid copepods, Bosmina longirostris and Daphnia galeata and D. hynlina. The population dynamics of Dapfinia had the strongest impact on all sestonic components. Daphnia maxima coincided with clearwater phases, and were negatively correlated with particulate organic carbon (POC), HNF and phytoplankton. Bacterial abundance was only slightly affected although daphnids were at times more important as bacterial consumers than HNF, as estimated from measured bacterial clearance rates. Other crustaceans (copepods, Bosmina) were probably of minor importance as grazers of bacteria and nanoplankton. 3. HNF abundance varied from 550 ml^?1 to more than 30000 ml^?1. HNF appeared to be suppressed by daphnids and reached highest densities when copepods dominated the metazooplankton. The variation in HNF abundance was not reflected in the concentration of heterotrophic bacteria, which fluctuated rather irregularly between 5 and 20 ± 10⁶ ml^?1. Long filamentous bacteria which were probably resistant to protozoan grazing, however, appeared parallel to the development of HNF. These bacterial cells, although small in number, could comprise more than 30% of the total bacterial biomass.     

Improved Method for Determining Bacterial Filtration Rates in Zooplankton

Filtration rates were determined for a natural population of zooplankton grazers (Bosmina longirostris [Müll.], Cyclops vicinus vicinus [Ulianine], Acanthodiaptomus denticornis [Wierz.], and Daphnia longispina [Müll.]) by using ³H-labeled bacteria as food for these organisms. There was a relationship between filtration rates of the major zooplankton grazers and the prevailing algal and bacterial composition in the lake water. Low filtration rates were obtained in the presence of colonial and filamentous cyanobacteria. The rapid process of bacterial adhesion to the external organs of grazers can result in an overestimation of filtration rates. By using the simple method presented here, filtration rates, with simultaneous correction for bacterial adhesion, can be quickly determined.     

Effects of fatty acid and phosphorus content of food on the growth, survival and reproduction ofDaphnia

1. Mixtures of microencapsulated lipids and Scenedesmus quadricauda grown at different degrees of P limitation were used as food for Daphnia galeata in two growth experiments. Thereby, food quality in terms of ω3-fatty acid (ω3-FA) or phosphorus (P) content could be assessed without interference from other factors.
2. ω3-highly unsaturated fatty acids (ω3-HUFA), given to Daphnia as fish oil or eicosapentaenoic acid (EPA) together with non P-limited algae, decreased the time to first reproduction. When fed fish oil, somatic growth and survival were also enhanced. Linolenic acid also decreased the time to first reproduction but to a lesser extent than EPA.
3. Food quality depended to a large extent on the degree of P limitation of Scenedesmus , which is consistent with P limitation of Daphnia. The overall impact of P was always larger than the effect of ω3-FA. Growth, survival and reproduction were elevated when Daphnia were fed non P-limited Scenedesmus compared to treatments with P-limited algae.
4. The relative importance of ω3-HUFA and P content in the food changed over a C : P gradient, with stronger effects of ω3-HUFA at low C : P ratios.     

Changes in the pelagic microbial food web due to artificial eutrophication

The effect of nutrient enrichment on the structure and carbon flow in the pelagic microbial food web was studied in mesocosm experiments using seawater from the northern Baltic Sea. The experiments included food webs of at least four trophic levels; (1) phytoplankton–bacteria, (2) flagellates, (3) ciliates and (4) mesozooplankton. In the enriched treatments high autotrophic growth rates were observed, followed by increased heterotrophic production. The largest growth increase was due to heterotrophic bacteria, indicating that the heterotrophic microbial food web was promoted. This was further supported by increased growth of heterotrophic flagellates and ciliates in the high nutrient treatments. The phytoplankton peak in the middle of the experiments was mainly due to an autotrophic nanoflagellate, Pyramimonas sp. At the end of the experiment, the proportion of heterotrophic organisms was higher in the nutrient enriched than in the nutrient-poor treatment, indicating increased predation control of primary producers. The proportion of potentially mixotrophic plankton, prymnesiophyceans, chrysophyceans and dinophyceans, were significantly higher in the nutrient-poor treatment. Furthermore, the results indicated that the food web efficiency, defined as mesozooplankton production per basal production (primary production + bacterial production − sedimentation), decreased with increasing nutrient status, possibly due to increasing loss processes in the food web. This could be explained by promotion of the heterotrophic microbial food web, causing more trophic levels and respiration steps in the food web.     

Good food versus bad food: the role of sterols and polyunsaturated fatty acids in determining growth and reproduction of <Emphasis Type="Italic">Daphnia magna</Emphasis>

The absence of dietary sterols and polyunsaturated fatty acids (PUFAs) has been shown to affect the performance of the freshwater herbivore Daphnia. Here, we compared somatic growth rates and clutch sizes of Daphnia magna reared on a diet of low food quality (Synechococcus elongatus) and of high food quality (Cryptomonas sp.) and investigated if and to what extent the absence of sterols or PUFAs in the cyanobacterium S. elongatus accounts for the observed differences in food quality. The supplementation of S. elongatus with cell-free lipid extracts (fatty acids, sterols, total lipids) obtained from the flagellate Cryptomonas sp. suggested that the superior food quality of Cryptomonas sp. is predominantly, but not completely, a combined effect of its sterol and PUFA composition. Our laboratory study suggests that somatic growth of D. magna feeding on S. elongatus is primarily constrained by the absence of sterols, whereas egg production is primarily limited by the absence of long chain PUFAs.     

Relationships between heterotrophic nanoflagellates and the demographic response of Daphnia longispina in a eutrophic lake with poor food quality conditions

1. Based on both field data and laboratory studies, the summer population of Daphnia longispina living in a stratified eutrophic lake was examined in relation to the abundance of algae, nanoflagellates and picocyanobacteria.
2. In early July, the Daphnia population replaced Bosmina and remained the dominant zooplankter during summer 1994. Its development in July was concomitant with an increase of edible algae but, despite the apparent abundance in available food, the Daphnia population decreased throughout August suggesting poor food conditions.
3. From mid-August to the beginning of September, the biomass of inedible phytoplankton was greater than that of the smaller, edible fraction. Consequently the average rate of increase, birth rates and fecundity of Daphnia remained low. Although the biomass of heterotrophic nanoflagellates was consistently low, the demographic parameters of Daphnia were correlated throughout this period with these protozoans.
4. Life table experiments run in the laboratory showed that epilimnetic food supported both the growth in length of individual Daphnia and an increase in fecundity, but metalimnetic food supported only individual growth. D. longispina probably failed to reproduce because of the abundance of detritus mixed with the heterotrophic nanoflagellates in the metalimnetic water at that period of the year. The vertical migration of Daphnia into these deeper layers could be caused by a predator avoidance mechanism.     

Predation-mediated shifts in size distribution of microbial biomass and activity during detritus decomposition

Many experimental studies on detritus decomposition revealed a comparable microbial succession after the addition of a substrate pulse: from small, freely suspended single bacteria at the beginning, to more complex and larger growth forms during a later stage, accompanied by the appearance of bacterivorous protists. We examined in three model experiments with different organic carbon sources whether this shift in bacterial size structure is linked to the grazing impact of bacterivores. In short‐term (8–10 d) microcosm experiments we added natural dissolved and particulate detritus (macrophyte leaves and leachate, dead phytoplankton cells) as an organic substrate source. By the use of size‐fractionated inocula and eucaryotic inhibitors we obtained treatments without protists, in which bacteria developed without predation. These were compared, by measurements of bacterial activity and microscopical analysis of bacterial size structure, to incubations in which either cultured heterotrophic nanoflagellates or a natural protist assemblage was included in the inoculum. The presence of bacterial grazers resulted in a 50–90% reduction of bacterial biomass compared to grazer‐free trials. The selective removal of freely suspended bacteria produced a very different relative composition of bacterial biomass: it became dominated by large, grazing‐resistant forms such as filaments and cells attached to particles or clustered in small aggregates. In grazer‐free treatments, bacterial biomass was always dominated (>80%) by free‐living, single bacterial cells. The time course of the bacterial development suggested different underlying mechanisms for the appearance of predation resistant filamentous and of aggregated or attached bacteria. As bacterial aggregates developed in approximately similar amounts with and without grazers no specific growth stimulation by protists could be detected. In contrast, concentrations of filamentous bacteria were 2–10 times higher in treatments with protists, thus indicating a stimulation of this growth form during enhanced grazing pressure. Measurements of ectoenzymatic activity and H‐leucine uptake indicated that microbial activity was also shifted to larger size fractions. In most cases more than 50% of bacterial activity in treatments with protists was associated with the size fraction>10 μm whereas this value was <2% without grazers. Grazing by protists also enhanced the specific activity of the bacterial assemblage which is in contrast to an assumed lower competitive ability of complex bacterial growth forms. The results imply that the selective force of bacterivory in nutrient‐rich environments changes the structure and possibly the function of aquatic bacteria and their position in the food web, making protist‐resistant bacteria more vulnerable to metazoan filter feeders and detritivores, and possibly also subject to sedimentation.     

Changes in food quality of the green alga Scenedesmus induced by Daphnia infochemicals: biochemical composition and morphology

1. The effects of Daphnia infochemicals on the morphology and biochemical composition of Scenedesmus were studied and subsequently their influence on Daphnia life history. Three species of Scenedesmus ( S. acutus, S. obliquus and S. subspicatus ) were tested for Daphnia -induced colony formation. Life history experiments were performed with Daphnia cucullata and D. magna feeding on unicellular or colonial S. acutus.
2. Colony formation was promoted when S. acutus and S. obliquus were exposed to filtered water from a D. magna culture. S. subspicatus did not form colonies when exposed to culture water, but showed some colony formation when exposed to live D. magna.
3. No clear differences were found in total lipid, protein or carbohydrate content between the unicellular and colonial ecomorphs of the three Scenedesmus species. However, fatty acid (FA) concentration and composition were changed. Total FA as a percentage of frozen dry weight (DW) were increased in colonies. The ratio of monounsaturated (MUFA) and polyunsaturated fatty acids (PUFA) and the ratio of ω3 : ω6 fatty acids decreased in colonies.
4. A short-term grazing experiment with three size classes of Scenedesmus revealed lower clearance rates for small Daphnia when feeding on large colonial Scenedesmus.
5. Life history parameters of both Daphnia species were influenced by food type. The intrinsic rate of population increase ( r) was significantly lower for daphnids feeding on colonies.
6. The negative influence of colonial Scenedesmus on population growth of Daphnia seems to be determined by algal morphology rather than biochemical composition.     

Daphnia performance on diets containing different combinations of high‐quality algae,heterotrophic bacteria,and allochthonous particulate organic matter

1. Filter‐feeding zooplankton in lakes feed on a mixture of phytoplankton, bacteria, and terrestrial particles and the proportions and nutritional value of these components can be highly variable. However, the extent to which food quality interacts with food quantity in affecting overall zooplankton performance is not yet fully resolved.
2. Here we performed laboratory feeding experiments to test how the performance of the unselective filter feeder Daphnia galeata was affected if various quantities of high‐quality food (the phytoplankton Rhodomonas) were diluted with low‐quality food such as heterotrophic bacteria (Pseudomonas) or terrestrial detritus particles (t‐POM) from the riparian zone of a boreal forest stream. We hypothesised: that increased proportions of bacteria and t‐POM in the diet will lead to decreased survival, somatic growth; and reproduction of Daphnia despite the presence of phytoplankton; that these effects are more pronounced for t‐POM than for heterotrophic bacteria; and that this response is stronger when phytoplankton availability is low.
3. Increasing the concentrations of Pseudomonas affected Daphnia survival, growth, and reproduction negatively when Rhodomonas was available at intermediate (0.37 mgC/L) and high (0.55 mgC/L) quantities. When Rhodomonas quantity was low (0.22 mgC/L), the addition of Pseudomonas generally resulted in better Daphnia performance except at very high concentrations of the bacterium relative to Rhodomonas. In contrast, the addition of t‐POM was detrimental for overall Daphnia performance at all Rhodomonas concentrations.
4. Daphnia performance was best described by a model including the interaction between food quality and quantity, with stronger negative effects on Daphnia when high‐quality food was supplemented with t‐POM than with Pseudomonas.
5. The results indicate that the ability of zooplankton to use low‐quality food is affected by the concurrent availability of high‐quality food. Furthermore, food sources that can be used but do not fulfil dietary requirements of grazers (e.g. bacteria), may still provide nutritional benefits as long as other complementary food components are available in sufficient quantities to compensate for biochemical deficiencies.
6. Therefore, we conclude that heterotrophic bacteria, but not peat layer t‐POM, can be an important component of zooplankton diets in boreal lakes, especially if the concentration of phytoplankton is low.

     

Comparative growth rates and yields of ciliates and heterotrophic dinoflagellates

Growth rates, ingestion rates and grazer yields (grazer volumeproduced/prey volume consumed) were measured for six protozoanspecies (ciliates: Favella sp., Strombidinopsis acuminatum,Uronema sp.; heterotrophic dinoflagellates: Amphidinium sp.,Gymnodinium sp., Noctiluca scintillans) in laboratory batchculture experiments. Comparative growth data indicate that theprymnesiophyte Isochrysis galbana, the prasinophyte Mantoniellasquamata, two cryptophyte species and several autotrophic dinoflagellatespecies were suitable foods for these grazers. When grown onoptimized diets at 13C, maximum ciliate growth rates (range0.77–1.01 day^–1 uniformly exceeded maximum heterotrophicdioflagellate growth rates (range 0.41–0.48 day^–1).A compilation of published data demonstrates that this growthrate difference persists across a range of ciliate and dinoflagellatetaxa and cell sizes. Comparison of volume-specific ingestionrates and yields for the six species studied here showed thatthere was no single explanation for this growth rate disparity.Heterotrophic dinoflagellates exhibited both low ingestion ratesand, in one case, low yields; ciliates were able to achievehigher growth rates via either higher ingestion rates or higheryields, depending on ciliate species. Volume yield increasedover time throughout the exponential growth phase in nearlyall experiments, suggesting variation in response to changingfood concentrations or long-term acclimation to culture conditions.Higher maximum ciliate growth rates mean that these grazershave the potential to exercise tighter control over incipientblooms of their prey than do heterotrophic dinoflagellates.     

Can phosphorus limitation inhibit dissolved organic carbon consumption in aquatic microbial food webs? A study of three food web structures in microcosms

Microcosms with three different food web structures and phosphorus (P) limited growth medium were used to study the interactions between P and organic carbon (C) fractions in pelagic food webs. The cultures were run with low dilution to allow the biological processes to determine the outcome. A double isotope technique was used to follow the C and P compartments. In all systems the primary production was P limited. The measured P:C ratios and the observed accumulation of degradable dissolved organic carbon (DOC) indicated that the growth of heterotrophic bacteria was also P limited. The presence of neither algal grazers nor flagellates feeding on bacteria altered the limitation pattern. A net loss of P from the bacterial fraction was observed after the bloom. Different strategies for nutrient aquisition and growth are proposed as mechanisms enabling simultaneous P limitation of algae and bacteria, and a concomitant accumulation of degradable DOC. The ability of the algae to grow with low P:C ratio keeps the regeneration of P through grazers low enough to cause sustained P limitation of both algae and bacteria. The grazers were important producers of DOC when present. This implies that the usual assumption of carbon limited bacterial growth may lead to wrong conclusions regarding the dynamics of plankton communities and the DOC pool.     

Effect of the addition of polyunsaturated fatty acids to the diet on the growth and fecundity ofDaphnia galeata

1. The importance of long-chain polyunsaturated fatty acids (PUFAs), in particular eicosapentaenoic acid (EPA), for the growth and development of Daphnia galeata (Sars) was tested using food types differing in PUFA and EPA contents.
2. Life history experiments of D. galeata fed with the cryptophyceans Rhodomonas lacustris (Pascher & Ruttner) and Cryptomonas pyrenoidifera (Gentler), and the green alga Scenedesmus acutus (Meyen), showed that both cryptophycean species were higher in quality than S. acutus.
3. Since the cryptomonads contained significant amounts of EPA while no EPA could be detected in Scenedesmus , tests were performed to ascertain whether EPA was responsible for the differences in food quality. Feeding daphnids a mixed diet of Scenedesmus and emulsion particles that were rich in EPA and DHA, resulted in a significant improvement in the intrinsic population growth rate. The initial difference in food quality between Scenedesmus and the cryptomonads was completely compensated for by addition of emulsion to the Scenedesmus food.
4. From the observed stimulatory effect of the addition PUFA to the daphnid diet, this study concludes that the presence of such long-chain PUFA improves food quality for daphnids.